The Genus Galadi: Three New Bandicoots (Marsupialia, Peramelemorphia) from Riversleigh’S Miocene Deposits, Northwestern Queensland, Australia K

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Journal of Vertebrate Paleontology Publication details, including instructions for authors and subscription information: http://www.tandfonline.com/loi/ujvp20 The genus Galadi: three new bandicoots (Marsupialia, Peramelemorphia) from Riversleigh’s Miocene deposits, northwestern Queensland, Australia K. J. Travouillon a b , Y. Gurovich b , M. Archer b , S. J. Hand b & J. Muirhead b a School of Earth Sciences, University of Queensland, St Lucia, Queensland, 4072, Australia b School of Biological, Earth and Environmental Sciences, University of New South Wales, New South Wales, 2052, Australia Version of record first published: 08 Jan 2013.

To cite this article: K. J. Travouillon , Y. Gurovich , M. Archer , S. J. Hand & J. Muirhead (2013): The genus Galadi: three new bandicoots (Marsupialia, Peramelemorphia) from Riversleigh’s Miocene deposits, northwestern Queensland, Australia, Journal of Vertebrate Paleontology, 33:1, 153-168 To link to this article: http://dx.doi.org/10.1080/02724634.2012.713416

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THE GENUS GALADI: THREE NEW BANDICOOTS (MARSUPIALIA, PERAMELEMORPHIA) FROM RIVERSLEIGH’S MIOCENE DEPOSITS, NORTHWESTERN QUEENSLAND, AUSTRALIA

K. J. TRAVOUILLON,*,1,2 Y. GUROVICH,2 M. ARCHER,2 S. J. HAND,2 and J. MUIRHEAD2 1School of Earth Sciences, University of Queensland, St Lucia, Queensland 4072, Australia, [email protected]; 2School of Biological, Earth and Environmental Sciences, University of New South Wales, New South Wales 2052, Australia

ABSTRACT—We describe three new bandicoot species of the genus Galadi from the Miocene of Riversleigh World Heritage Area in northern Australia. The ﬁrst species, which is represented by a complete dentary and several isolated upper molars, is restricted to Riversleigh’s Faunal Zone B. It is the largest bandicoot as yet known from Riversleigh. The second species is represented by 19 specimens, including a partial skull and several maxillae and dentaries, from Riversleigh’s Faunal Zone C. Several features distinguishing this species from the similarly sized type species G. speciosus are of interest, notably the presence of larger maxillopalatine fenestrae and additional maxillary fenestrae, incomplete centrocrista on all upper molars, a more complete posterior cingulum on upper molars, and higher degree of dental wear, which together suggest a more omnivorous diet. The third species is represented by a single maxillary, which exhibits a quite different combination of dental features compared with other Galadi species. All Galadi species appear to be restricted to Riversleigh’s Faunal Zones B and C, which are interpreted to be early and middle Miocene in age, respectively, with rainforest habitats persisting throughout. SUPPLEMENTAL DATA—Supplemental materials are available for this article for free at www.tandfonline.com/UJVP

INTRODUCTION described from Riversleigh Oligo-Miocene sites (Muirhead, 2000; Archer et al., 2006) and eight from the Etadunna and Wipajiri Peramelemorphia (bandicoots and bilbies) is an extant or- formations of South Australia (Woodburne et al., 1993). der of marsupials from Australia, New Guinea, and surround- Here we describe three new bandicoot species from the ing islands. It consists of the two extant families of Peramel- Miocene of Riversleigh that appear to be referable to Galadi. The idae: Thylacomyidae and the recently extinct Chaeropodidae three species are represented by numerous specimens, including (which contains Chaeropus ecaudatus, the pig-footed bandicoot). partial cranial remains and isolated maxillae and dentaries. We Eight extant Australian species are referred to four genera (Per- assess the phylogenetic relationships of these three taxa within ameles, Isoodon, Macrotis,andEchymipera), with at least 13 Peramelemorphia. We also discuss their paleoecology and docu- New Guinean species (in the genera Echymipera, Peroryctes, Mi- ment the temporal distribution of species of the genus Galadi. croperoryctes,andRhynchomeles; see also Aplin et al., 2010). Currently, nine fossil species of peramelemorphians are known from the late Oligocene to the Pleistocene of Australia; six of these are attributed to modern families and two are referred to MATERIALS AND METHODS the extinct family Yaralidae (Muirhead, 2000).

Downloaded by [58.165.130.223] at 03:54 06 February 2013 All specimens were collected from the Riversleigh WHA, Galadi is the second genus of fossil bandicoots recognized northwestern Queensland, between 1983 and 2010, and recov- from the Oligo-Miocene of Riversleigh World Heritage Area ered by acid processing of limestone blocks. Biostratigraphic (WHA), northwestern Queensland, Australia. It is currently nomenclature follows Arena (2004) and Travouillon et al. (2006). known only by the type species Galadi speciosus, represented by Institutional Abbreviations—AR, Specimens temporarily held an exceptionally well-preserved skull and a few dentary remains in collections at the University of New South Wales, Sydney; (Travouillon et al., 2010). QMF, Queensland Museum fossil collection, Brisbane. Galadi speciosus exhibits a mixture of plesiomorphic and apo- Anatomical Abbreviations—Dental terminology follows morphic dental and cranial characters, with the relationship be- Archer (1976), Muirhead and Filan (1995), Voss and Jansa tween Galadi speciosus and Yarala burchfieldi (the first per- (2003), and Travouillon et al. (2010) and with molar loci homol- amelemorphian described from the same Oligo-Miocene deposits ogy following Luckett (1993). Cranial anatomy follows Muirhead of Riversleigh by Muirhead 2000) being unresolved (Travouil- (1994, 2000), Voss and Jansa (2003), Wible (2003), Beck et al. lon et al., 2010). However, Galadi speciosus appears to lie out- (2008a), and Travouillon et al. (2010). Peramelemorphian sys- side crown group Peramelemorphia but closer to the crown group tematics follows Groves and Flannery (1990), Muirhead (1994, than Yarala burchfieldi, and for this reason, was not referred to 2000), Muirhead and Filan (1995), and Travouillon et al. (2010). the family Yaralidae (Travoullion et al., 2010). Higher-level marsupial systematics follows Aplin and Archer A total of three species of fossil bandicoots have been de- (1987). scribed from the Oligo-Miocene of Australia: Yarala burchfieldi and Galadi speciosus from the Riversleigh WHA; and Yarala kida Schwartz, 2006, from the late Oligocene Kangaroo Well Lo- Phylogenetic Analysis cal Fauna. At least six additional bandicoot species are yet to be We analyzed the matrix from Travouillon et al. (2010) and made a number of changes. The original matrix contained 51 qualitative morphological characters (28 dental and 23 cranial). *Corresponding author. 153 154 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 33, NO. 1, 2013

We made the following changes and additions, resulting in a total Revised Generic Diagnosis—Species of Galadi differ from of 74 characters: all other bandicoots in the following combination of features: short and robust snout; upper molar crown margins rounded, 1. Character 15, ‘morphology of the centrocrista,’ was split into short and wide, lower molar crown anteroposteriorly reduced; two characters, to distinguish between the morphology of the anterior cingula on all upper molars; large entoconid, blade-like centrocrista of M1 and M2 from that of M3. with low anteriorly directed preentocristid; small metaconular 2. We added 14 characters from Voss and Jansa (2009; characters hypocone; parastylar region of M1 less blade-like than in modern 52, 55, 57, 59, 82, 85, 89, 93, 96, 97, 114, 116, 120, and 125). peramelemorphians; and large size. Upper and lower premolars 3. We added four characters from Aplin et al. (2010; characters are large and wide. Anterior cusps of lower premolars are absent 3, 6, and 8 from page 22 and character 11 from page 17) or minute. 4. We added four new dental characters (shape of I2–4, shape Five incisors without separating diastemata, nasals extend pos- of upper canine, lower molar crown height, and presence of teriorly past the anterior region of the orbit; short infraorbital metaconule). canal; no lachrymal orbital rim; moderate nasolabial/antorbital All new characters were left unordered, leaving the original 23 or- fossa; large orbitosphenoid; alisphenoid and parietal contact; un- dered characters in all analyses (see online Supplementary Data specialized sphenorbital fissure and foramen rotundum; one pair 1 and 2 for a complete list of characters) of incisive foramina and one pair of maxillopalatine fenestrae; We included the 21 peramelemorphian species surveyed by large primary foramen ovale bordered by alisphenoid and pet- Travouillon et al. (2010) and 12 additional taxa, of which four rosal; no secondary foramen ovale; small, uninflated alisphenoid are extant (Peroryctes broadbenti, Microperoryctes papuensis, tympanic wing; small knob-like rostral tympanic process of the M. ornata, Echymipera davidi) and eight are fossil taxa from petrosal; uninflated epitympanic recess; and very weakly defined the Pleistocene (Perameles sobbei; Price 2002), Pliocene (Ischn- squamosal epitympanic sinus. odon australis, Stirton, 1955; Numbigilga ernielundeliusi,Beck, Archer, Godthelp, Mackness, Hand, and Muirhead, 2008a; cf. GALADI GRANDIS, sp. nov. Peroryctes tedfordi and cf. Peroryctes. sp., Turnbull et al., 2003), (Figs. 1–4) and Miocene (the three new taxa described here). Following Travouillon et al. (2010), we used the following Specific Diagnosis—Galadi grandis differs from the type taxa as outgroups: the early Eocene australidelphian Djarthia species G. speciosus in the following combination of features: murgonensis Godthelp, Wroe, and Archer, 1999; the early having a complete centrocrista on all upper molars that does Miocene dasyurid Barinya wangala Wroe, 1999; and the middle not breech the ectoloph; stylar cusp E present on M1 and M2; Miocene dasyuromorphian Mutpuracinus archibaldi Murray and smaller metaconular hypocone on upper molars making the over- Megirian, 2006. all shape more triangular; laterally compressed lower canine; m1 paraconid is more buccally positioned, curving the anterior part of the tooth between the paraconid and metaconid in occlusal Parsimony Analysis view; and the dentary is longer and less robust at the anterior Parsimony analysis of the matrix (see online Supplementary end. Size large (∼1.5–2 kg compared with ∼900 g). Data) was carried out using PAUP∗ 4.0b10 (Swofford, 2002), fol- Specific Etymology—‘Grandis’, Latin word meaning grand, in lowing Travouillon et al. (2010), using an initial search comprising reference to its large size (being the largest bandicoot so far 1000 heuristic replicates, saving 10 trees per replicate, followed known from the Riversleigh WHA). by a second heuristic search within the trees saved from the first Holotype—QM F23394, left dentary with c to m4 (Figs. 1–3). stage. Multiple most parsimonious trees were summarized using Paratypes—QM F23395, isolated LM1; QM F23396, isolated strict consensus. RM2; and QM F23397, isolated LM3 (Fig. 4). Bootstrap values were calculated using 100 bootstrap repli- Referred Material—Camel Sputum Site: QM F54575, iso- cates; each bootstrap replicate itself comprised 10 random addi- lated right M2; QM F54576, isolated right m2; QM F54577, tion sequence replicates. isolated right m3; QM F54578, isolated left m3; QM F54579, We performed two analyses, one containing all taxa, and a sec- isolated left m1. Creaser’s Rampart Site: QM F30788, left den-

Downloaded by [58.165.130.223] at 03:54 06 February 2013 ond in which we removed all taxa that were represented by max- tary fragment with unerupted p3 and loose m3. Mike’s Menagerie illa or dentary only. If one taxon is known only from a maxilla and Site: QM F56066, isolated left m2; QM F56065, isolated left m3; another is known only from a dentary, a phylogenetic analysis can QM F56067, isolated right m3. Ross Scott-Orr (RSO) Site: QM never reconstruct them in a clade (even if they are in fact closely F56062, isolated right M2. Wayne’s Wok Site: QM F54574, iso- related) because they have no characters scored in common, so lated left M2; QM F56063, isolated right M2. there is nothing that can link them. The taxa we removed from Type Locality—The holotype and paratypes are from Camel the second analysis were Galadi adversus, Perameles sobbei, Is- Sputum Site, Godthelp Hill, Riversleigh WHA, northwestern chnodon australis, Numbigilga ernielundeliusi,cf. Peroryctes ted- Queensland, Australia. fordi,andcf. Peroryctes sp. Age and Stratigraphy—These sites are part of Riversleigh WHA, northwestern Queensland, and contain terrestrial carbon- SYSTEMATIC PALEONTOLOGY ates that appear to represent vadose cave deposits (Arena, 2004). They are part of Faunal Zone B, estimated to be approximately Order PERAMELEMORPHIA (Kirsch, 1968) Aplin and early Miocene in age (Travouillon et al., 2006, 2011). Archer, 1987 Superfamily incertae sedis Family incertae sedis Description Genus GALADI Travouillon, Gurovich, Beck, and Muirhead, Dentary—Dorsal surface of the ramus remains at constant 2010 height (Fig. 1), with depth of ramus greatest below m3. The Type Species—Galadi speciosus Travouillon, Gurovich, Beck, symphysis extends posteriorly to below the posterior root of p2. and Muirhead, 2010. Two mental foramina are present: the anterior foramen is below Remarks—A nearly complete skull, with associated femur and p1, the posterior foramen below m1. The largest diastema occurs pelvis (QM F54568), of Galadi speciosus is now known from In- between c and p1 and a smaller one between p1 and p2. The abeyance Site, Faunal Zone B, Riversleigh WHA, northwestern coronoid process extends from the dentary at approximately Queensland. 95 degrees. TRAVOUILLON ET AL.—FOSSIL BANDICOOTS FROM AUSTRALIA 155

FIGURE 1. Galadi grandis, sp. nov., mandible (holotype QM F23394). A, buccal view of left mandible; B, lingual view of left mandible. Abbrevi- ations: ang, angular process; c, lower canine; coc, coronoid crest; con, mandibular condyle; m1–m4, lower molar 1 to lower molar 4; maf, masseteric fossa; manf, mandibular foramen; mas, mandibular symphysis; mf, mental foramen; p1–p3, lower premolar 1 to premolar 3. Scale bar equals 2 cm. Downloaded by [58.165.130.223] at 03:54 06 February 2013

Lower Dentition—The canine is laterally compressed (Fig. 1). cal to the metaconid, curving the anterior part of the tooth be- It is broken at its tip but would have been taller than the mo- tween the paraconid and metaconid in occlusal view. The talonid lars. It extends from the dentary at an oblique angle and is not is wider than the trigonid. The entoconid, oval in shape in occlusal strongly curved. The posterior roots of the premolars are thicker view, is posterolingually positioned relative to the metaconid and than the anterior roots (Fig. 2). Only remnants of an anterior cusp joined to the base of the metaconid by a low preentocristid. The are visible on p1 to p3. The anterior margin of the main cusp is hypoconulid is not raised but extends posteriorly; it is not joined rounded on p1 to p3. The main cusps of p2 and p3 are anteriorly to the entoconid but connects to the hypoconid by a deflected positioned above the gap between the roots, and lingually so that posthypocristid. The hypoconid lies directly buccal to the ento- the buccal surface bulges in occlusal view. The posterior cusp in- conid. The cristid obliqua is slightly concave, terminating in the creases in size from p1 to p3 (Fig. 2). The posterior crest curves buccal half of the tooth on the posterolingual flank of the proto- from the lingual main cusp to terminate at the middle of the pos- conid. A low ridge of enamel joins the protoconid to hypoconid. terior edge of the crown. No enamel ridge is present around the No posterior cingulid is present. posterior margin of p1 but is present on p2 and enlarged on p3 on Morphology of m2 is similar to that of m1 except as fol- both lingual and buccal surfaces. A crest runs around this ridge lows. The paraconid is directly anterior to the metaconid. The on p3. The crown and central cusp increase in size from p1 to p3. paracristid is longer. The distance between the metaconid and On m1, the protoconid and metaconid are closer together protoconid is greater but less than that between the para- than either is to the paraconid (Fig. 3). The paraconid is much conid and protoconid. In the trigonid, the smallest dimension lower on the crown, lower than the entoconid. No anterior cin- is between the paraconid and the metaconid. The anterior cingulid is present. The metaconid is posterolingually positioned gulid is wide and continuous from the tip to the anterobuccal with respect to the protoconid. The paraconid lies anterobuc- flank of the protoconid. The entoconid lies directly posterior to 156 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 33, NO. 1, 2013

FIGURE 3. Galadi grandis, sp. nov., lower molars (holotype QM F23394). A, occlusal view; B, buccal view. Abbreviations: acd,ante- FIGURE 2. Galadi grandis, sp. nov., lower premolars (holotype QM rior cingulid; co, crista obliqua; end, entoconid; hyd, hypoconid; hyld, F23394). A, occlusal view; B, buccal view. Abbreviations: p1–p3, lower hypoconulid; m1–m4, lower molar 1 to lower molar 4; mecd, metacristid; premolar 1 to premolar 3. Scale bar equals 2 mm. med, metaconid; pacd, paracristid; pad, paraconid; ped, preentocristid;

Downloaded by [58.165.130.223] at 03:54 06 February 2013 phd, posthypocristid; prd, protoconid. Scale bar equals 2 mm.

the metaconid. The cristid obliqua terminates midway between the metaconid and protoconid. The angle formed by crests of lum forms a continuous shelf from the anterolabial corner of the hypoconid is reduced. the tooth to the protocone via a connection with the prepro- Morphology of m3 is similar to that of m2 except as fol- tocrista. Stylar cusp B is posterobuccal to the paracone. In oc- lows. The protoconid lies directly anterior to the hypoconid. The clusal view, the preparacrista curves posteriorly to stylar cusp B paracristid and metacristid are longer. The talonid and trigonid from the paracone. There is no connection of stylar cusp A to the are equivalent in width. The cristid obliqua is concave at the an- paracone. Stylar cusp A is lower than the parastylar shelf. The terior end and terminates on the lingual half of the crown. stylar shelf crest is continuous from the preparacrista, through Morphology of m4 is similar to that of m3 except as follows. the stylar cusp B to the posterobuccal corner of the crown. Sty- The trigonid basin is anteroposteriorly widened. The talonid is re- lar cusp D is longer than wide and directly buccal to the meta- duced, but two cusps are retained: hypoconid and entoconid. The cone. Stylar cusp E is small but prominent. The centrocrista crista obliqua and preentocristid meet just anteriorly to the mid- does not breech the ectoloph. The postprotocrista runs paral- point between the entoconid and hypoconid. The posthypocristid lel to the stylar shelf before curving around the base of the meets the postentocristid posteriorly to the hypoconid. metaconular hypocone. It then thins along the posterior ﬂank of Upper Dentition—Occlusal shape of M1 rounded to triangu- the metacone to terminate as the posterior cingulum just prior lar (Fig. 4). The only specimen is worn. Stylar cusp D is the the crown’s posterior corner. The anterior cingulum continues tallest cusp, followed by stylar cusp B, metacone, stylar cusp E, through the protocone to the posterior cingulum rounding the lin- paracone, stylar cusp A (referred to as anterior cingulum tip gual edge of crown. The buccal ﬂank of the crown is bulbous and in Travouillon et al., 2010), and protocone. The anterior cingu- rounded. TRAVOUILLON ET AL.—FOSSIL BANDICOOTS FROM AUSTRALIA 157

The width is greater than length. Stylar cusp C is small and lies on the anterior flank of stylar cusp D. Stylar cusp E is absent. No continuous crest is present from stylar cusp D to the posterobuccal corner of the crown. The posterior crest of stylar cusp B terminates prior to stylar cusp C. The metaconular hypocone is present but small. The postprotocrista terminates at the posterolingual base of the metaconular hypocone but a ridge runs along the posterolingual flank to the posterior corner of the crown. Measurements of the upper dentition of G. grandis are pre- sented in Table 1 and those of the lower dentition in Table 2. GALADI AMPLUS, sp. nov. (Figs. 5–9) Specific Diagnosis—Galadi amplus differs from G. grandis and G. speciosus in the following combination of features: upper and lower third premolar are raised, exposing the anterior roots and reclining toward M1; P3 is wider with a wider lingual shelf extending more anteriorly; postparacrista and premetacrista do not connect on any upper molar to form the centrocrista; stylar cusp E, if present, only on M3; stylar cusp B absent on M4; small anterior cingulid present on m1; distance between paraconid and metaconid is relatively shorter on all molars; lower molar length is relatively shorter but wider at the talonid; large conical entoconid present on m4; infraorbital foramen lies above anterior rather than the posterior root of P3; sagittal crest extends further anteriorly, up to half the length of the frontal; wider and longer canine alveoli with bone thickening; posterior opening of the in- FIGURE 4. Galadi grandis, sp. nov., upper molars (top to bottom, ternal nares wider; and palate more fenestrated than in G. specio- paratypes QM F23395, QM F23396, and QM F23397). A, occlusal view; sus, with larger maxillopalatine fenestrae and additional pair of B, lingual view. Abbreviations: ac, anterior cingulum; cc, centrocrista; maxillary fenestrae. ect, ectoloph; M1–M3, upper molar 1 to upper molar 3; mcl, meta- Specific Etymology—The species epithet ‘amplus’, Latin, conular hypocone; me, metacone; pa, paracone; pc, posterior cingu- means ‘large’, in reference to large maxillopalatine fenestrae. lum; pmc, premetacrista; pomc, postmetacrista; popc, postparacrista; Holotype—QM F23398, partial skull including fragments of ppc, preparacrista; pr, protocone; prprcr, preprotocrista; psprcr, postprotocrista; St A, St B, St C, St D, St E, stylar cusps A, B, C, D, and E. Scale nasal, premaxilla, maxilla, frontal, jugal, palatine, lacrimal, al- bar equals 2 mm. isphenoid, squamosal, parietal, and basisphenoid. Contains broken M1 and complete but worn M2–4 (Fig. 5). Paratypes—QM F23399, right maxillary fragment with C, P3, M2–3 (Figs. 5, 7); QM F56072, right dentary with c, p1–3, m1–4 (Figs. 6–8); QMF23401, left maxillary fragment with M2–4; Morphology of M2 is similar to that of M1 except as follows. (Fig. 9); QM F56074, left maxilla with M1–2, M4 (Fig. 9). The occlusal outline of the crown is more angular. Stylar cusp B Referred Material—Cleft of Ages Site: QM F56068, left den- is as large as stylar cusp D. Stylar cusp A is on the parastylar shelf tary with m1–4. Gag Site: QMF23400, right maxillary fragment and has a crest joining to the preparacrista. The anterior cingulum with M1–3; QMF23402, right dentary fragment with p3, m2; is continuous from the base of stylar cusp A to the preprotocrista QMF23403, right dentary with p3–m3; QMF24226, right dentary after forming a weak notch with the anterolabial corner of the fragment with p3, m3–4; QM F56069, right maxilla with m2–3; tooth. An anterior crest links stylar cusp B to the preparacrista Downloaded by [58.165.130.223] at 03:54 06 February 2013 QM F56075, right dentary with p2–3, m1–2; QM F56064, left and in occlusal view bends towards stylar cusp A. No posterior dentary with m3–4; QMF 23404, left dentary with c, p2, m2–3; crest links stylar cusp B to stylar cusp D. The posterior cingulum QM F56070, right dentary with m3–4. Henk’s Hollow Site: QM is thinner than on M1. The protocone is more angular. The post- F56076, isolated left M3. Last Minute Site: QM F56071, right den- protocrista does not run parallel to the stylar shelf; the protocone tary with m2–3; QM F56073, left dentary with m3–4. Wang Site: is the most lingual portion of the crown. QMF 36308, left maxilla with M3–4. M3 morphology is similar to that of M2 except as follows. The Type Locality—The holotype QM F23398 and paratypes QM crown is much more triangular and angular in occlusal outline. F23399 and QMF23401 are from Gag Site, paratype QM F56072

TABLE 1. Measurements (in mm) of the upper dentition of type and referred material of Galadi grandis.

P1 P2 P3 M1 M2 M3 M4 Specimen Riversleigh locality L W L W L W L W L W L W L W QM F23395 Camel Sputum Site 5.37 3.92 QM F23396 Camel Sputum Site 4.82 4.35 QM F23397 Camel Sputum Site 4.75 4.53 QM F54575 Camel Sputum Site 4.83 4.4 QM F56062 Ross Scott-Orr Site 4.73 3.95 QM F54574 Wayne’s Wok Site 4.67 4.05 QM F56063 Wayne’s Wok Site 4.64 3.62

Abbreviations: L, anteroposterior length; M,molar;P, premolar; W, lingual-buccal width. 158 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 33, NO. 1, 2013

TABLE 2. Measurements (in mm) of the lower dentition of type and referred material of Galadi grandis.

p1 p2 p3 m1 m2 m3 m4 Specimen Riversleigh locality L W L W L W L AW PW L AW PW L AW PW L AW PW QM F23394 Camel Sputum Site 2.28 1.06 3.28 1.62 3.75 2.23 4.42 2.15 2.42 4.44 2.45 2.72 4.55 2.55 2.55 4.72.45 1.64 QM F54576 Camel Sputum Site 4.91 2.61 3.01 QM F54577 Camel Sputum Site 4.78 2.78 2.87 QM F54578 Camel Sputum Site 4.72 2.66 2.77 QM F54579 Camel Sputum Site 4.88 2.38 2.64 QM F30788 Creaser’s Rampart Site 4.52.55 2.72 QM F56066 Mike’s Menagerie Site 4.84 2.53 2.87 QM F56065 Mike’s Menagerie Site 4.99 2.87 2.85 QM F56067 Mike’s Menagerie Site 4.98 2.75 2.79

Abbreviations: AW, anterior width; L, anteroposterior length; m,molar;p, premolar; PW, posterior width.

is from Henk’s Hollow Site, and paratype QM F56074 is from has strong curvature around the canine root, more so than in Last Minute Site. These sites are on Gag Plateau, Riversleigh G. speciosus and the bone surrounding the canine alveoli is WHA, northwestern Queensland, Australia. thicker. Anteriorly, in dorsal/ventral view, the rostral end of Age and Stratigraphy—The sites are part of Riversleigh WHA, the nasal is rounded. Posteriorly, the nasal-frontal suture is northwestern Queensland, and contain terrestrial carbonates that saw-edged and is posterior to the orbit. A small pointed tongue appear to represent tufaceous deposits (Arena, 2004). They are of frontal extends between the nasal and maxilla. The premaxilla part of Faunal Zone C, estimated to be approximately middle is thin, extending to above P2. The most posterior end of the Miocene in age (Travouillon et al., 2006, 2011). premaxilla terminates within the maxilla and ends half way through the length of the nasal. The infraorbital foramen is located directly above the anterior root of P3, as opposed to the Description posterior root in G. speciosus. The lateral posterior surface of this Facial Skeleton—The skull is poorly preserved (Fig. 5). It foramen inclines obliquely. The distance between the infraorbital has short, broad snout, comparable to G. speciosus. The snout foramen and the lacrimal foramen is short, but longer than in is wider than tall and curves ventrally to the tip. The maxilla G. speciosus. The nasolabial/antorbital fossa is shallow compared Downloaded by [58.165.130.223] at 03:54 06 February 2013

FIGURE 5. Galadi amplus, sp. nov., skull (holotype QM F23398) and maxillary (paratypes QM F23399). A, dorsal view of skull; B, ventral view of skull; C, right lateral view of skull; D, left lateral view of skull; E, right lateral view of maxillary. Abbreviations: aofo, anterorbital fossa; as, alisphenoid; C, upper canine; fr, frontal; fro, foramen rotundum; inf, incisive foramen; iof, infraorbital foramen; ju, jugal; lac,lacrimal;lacf, lacrimal foramen; M2–M4, upper molar 2 to upper molar 4; mxf, maxillary fenestra; mp, maxillopalatine fenestra; mx, maxilla; na, nasal; pa, parietal; pal, palatine; plpf, posterolateral palatal foramen; pmx, premaxilla; ps, presphenoid; pt, pterygoid; P3, upper premolar 3; sg, saggital crest; sof, sphenorbital ﬁssure. Scale bar equals 2 cm. TRAVOUILLON ET AL.—FOSSIL BANDICOOTS FROM AUSTRALIA 159

FIGURE 6. Galadi amplus, sp. nov., mandible (paratype QM F56072). A, buccal view of right mandible; B, lingual view of right mandible. Abbre- viations: c, lower canine; m1–m4, lower molar 1 to lower molar 4; maf, masseteric fossa; manf, mandibular foramen; mas, mandibular symphysis; mf, mental foramen; p1–p3, lower premolar 1 to premolar 3. Scale bar equals 2 cm.

with modern peramelemorphians and despite the jugal being Cranial Floor—The posterior opening of the internal nares is

Downloaded by [58.165.130.223] at 03:54 06 February 2013 poorly preserved, the maxilla-jugal suture is visible as a narrow large, wider than in G. speciosus. Contact between the presphe- V-shaped suture. The maxilla drops ventrally in the region of the noid and pterygoid is unclear (Fig. 5). The basicranial region is molars. Very little of the lacrimal extends onto the facial region absent. and surrounds anteriorly the lacrimal foramen, just anterior to Dentary—Two mental foramina are present, the first lies be- the orbit. No distinct lacrimal tubercle or crests are present. neath the anterior root of p1, the second beneath the anterior Neurocranium—Lacrimal sutures are poorly preserved. In root of m1 (Fig. 6). The dentary is overall curved anteroposteri- dorsal view, the frontal is widest anterior to the orbit, then nar- orly. The posterior part of the dentary is poorly preserved. The rows, and widens again at its posterior end (Fig. 5). A high sagit- base of the coronoid process extends from the ramus at 100 de- tal crest is present in the posterior region of the frontal. Very grees. little of the parietal is preserved. Contact with the alisphenoid is Lower Dentition—No lower incisors are preserved. The in- not visible. The maxillary foramen is large and wholly bounded cisor region has alveoli for three single-rooted incisors. As in G. by the maxilla. The sphenopalatine foramen is large and round. speciosus, the alveolus of i2 is raised to a level above that of i1. The posterolateral palatal foramen is poorly preserved but would The canine is large and laterally flattened (Fig. 6). The canine have been larger than in G. speciosus. The sphenorbital fissure is is tall, well above molars and premolars. The canine and p1 are preserved; the foramen rotundum is lacking only its lateral wall. separated by a short diastema. Palate—Only the posterior end of one incisive foramen is pre- The morphology of p1, p2, and p3 is very similar, only differing served (Fig. 5). The palate region of the premolars has a pair of in size, which increases posteriorly (Figs. 6–7A). All premolars small irregularly shaped maxillary fenestrae, extending from P1 have only two cusps, with the major cusp anteriorly positioned to P2. In the molar region, a pair of large maxillopalatine fenes- above the anterior root. The posterior cusp is directly positioned trae is present, oval in shape with a thick dividing septum. They posterior to the major cusp, and a crest joins the two cusps. This extend from level with the posterior root of P3 to level with the crest on p2 and p3 is heavily worn posteriorly. No cusp or crest anterior root of M4 lies anterior to the main cusp. The premolar crowns are raised 160 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 33, NO. 1, 2013

FIGURE 7. Galadi amplus, sp. nov., lower premolars (paratype QM F56072) and upper third premolar (paratypes QM F23399). A, occlusal view of lower premolars; B, occlusal view of upper third premolar. Ab- breviations: p1–p3, lower premolar 1 to premolar 3; P3, upper third premolar. Scale bar equals 2 mm. FIGURE 8. Galadi amplus, sp. nov., lower molars (paratype QM

Downloaded by [58.165.130.223] at 03:54 06 February 2013 F56072). A, occlusal view; B, buccal view. Abbreviations: acd,anterior cingulid; co, crista obliqua; end, entoconid; hyd, hypoconid; hyld, hypoconulid; m1–m4, lower molar 1 to lower molar 4; mecd, metacristid; above the dentary by extension of the roots. The p1 leans anteri- med, metaconid; pacd, paracristid; pad, paraconid; ped, preentocristid; orly, whereas p2 and p3 lean posteriorly. phd, posthypocristid; prd, protoconid. Scale bar equals 2 mm. The anterior cingulid of m1 is short (Fig. 8). The paraconid is low. The metaconid is taller than the protoconid. The metaconid and protoconid are closer to each other than either is to the paraconid. The metaconid is almost directly lingual to the protoconid. widens at the base of the protoconid. The paraconid is lower than The entoconid is larger than the paraconid and taller than the the protoconid. The cristid obliqua terminates below the center hypoconid. The entoconid is directly posterior to the metaconid. of the metacristid. There is no connection of the preentocristid The hypoconulid is low, posterobuccal to the entoconid, and con- to the trigonid. The preentocristid is short, anteriorly directed, nected to the hypoconid by the posthypocristid. The cristid obli- terminating at the anterior base of the entoconid. The worn qua is straight, terminating on the buccal side of the trigonid. entoconid is wider posteriorly than anteriorly, almost tear- The talonid is wider than the trigonid. The hypoconid is directly shaped. The trigonid and talonid are wider than on m1 but the buccal to the entoconid. The posthypocristid is parallel to the trigonid remains narrower than the talonid. metacristid and is almost perpendicular to the tooth row. Morphology of m3 is similar to that of m2 except as follows The morphology of m2 (Fig. 8) is similar to that of m1 except (Fig. 8). The protoconid is more buccally oriented, lengthening as follows. The paraconid is closer to the metaconid than either the paracristid and metacristid. The anterior portion of the ante- is to the protoconid. The paraconid is thin compared with the rior cingulid is curved laterally. metaconid. The metacristid is perpendicular to the tooth row and The morphology of m4 is similar to that of m3 except as fol- parallel to the posthypocristid. The anterior cingulid is long and lows (Fig. 8). The anterior cingulid is wider and straighter. The TRAVOUILLON ET AL.—FOSSIL BANDICOOTS FROM AUSTRALIA 161 Downloaded by [58.165.130.223] at 03:54 06 February 2013

FIGURE 9. Galadi amplus, sp. nov., upper molars (top to bottom, paratypes QM F56074 and QM F23401). A, occlusal view; B, lingual view. Abbre- viations: ac, anterior cingulum; cc, centrocrista; ect, ectoloph; M1–M3, upper molar 1 to upper molar 3; mcl, metaconular hypocone; me, metacone; pa, paracone; pc, posterior cingulum; pmc, premetacrista; pomc, postmetacrista; popc, postparacrista; ppc, preparacrista; pr, protocone; prprcr,preprotocrista; psprcr, postprotocrista; St A, St B, St C, St D, St E, stylar cusps A, B, C, D, and E. Scale bar equals 2 mm. 162 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 33, NO. 1, 2013

metaconid is larger and equidistant from paraconid and protoconid. The protoconid is the tallest cusp. The talonid is reduced in both width and length. The hypoconid is more lingually positioned. The entoconid is a conical cusp, taller than the hypoconid. No distinct hypoconulid is present. The cristid obliqua terminates on the lingual side of the trigonid. Upper Dentition—The incisor region is not preserved. The canine is strongly curved and large (Fig. 5E). This broken specimen shows no indication of any immediate tapering, suggesting a long length. It is laterally flattened. A short diastema is present between canine and alveoli for P1. P1 or P2 are not preserved. The alveoli of P1 are aligned parallel to the remaining tooth row. A large diastema is present between P1 and P2. The alveoli of P2 are the same as for P1 except the posterior is slightly larger than the anterior. No diastema separates P2 from P3. The crown of P3 is raised well above the maxilla with the roots exposed. The anterior root is raised further than the posterior root so that the tooth reclines against M1. The posterior root is wider than the anterior root. The main cusp is centrally located on the crown (Fig. 7B). A prominent crest joins the main cusp to the posterior-most corner of the crown. No anterior cusp or crest is present. The anterior margin is rounded. A cingulum is present on the posterobuccal surface and is continuous posteriorly to the anterolingual surface. Lingual development of the cingulum is greater than buc- FIGURE 10. Galadi adversus, sp. nov., upper molars (holotype QM F16911). A, occlusal view; B, lingual view. Abbreviations: ac,anterior cally with increased width lingual to the main cusp. cingulum; cc, centrocrista; ect, ectoloph; M1–M3, upper molar 1 to up- In occlusal view, M1 is rounded on all margins (Fig. 9). Its per molar 3; mcl, metaconular hypocone; me, metacone; pa, paracone; buccal length is greatest. Stylar cusp D is the tallest cusp, fol- pc, posterior cingulum; pmc, premetacrista; pomc, postmetacrista; popc, lowed in decreasing height by the posterolabial corner of the postparacrista; ppc, preparacrista; pr, protocone; prprcr, preprotocrista; tooth (referred to as the metastylar tip in Travouillon et al., 2010), psprcr, postprotocrista; St A, St B, St C, St D, St E, stylar cusps A, B, C, stylar cusp B, metacone, stylar cusp A, paracone, metaconular D, and E. Scale bar equals 2 mm. hypocone, and protocone. The preparacrista runs buccally from the paracone, then curves posteriorly to join stylar cusp B, continues through stylar cusp D, and ends at the posterolabial corner of the tooth. The postparacrista is straight and obliquely ori- much smaller than on M2. The postprotocrista is therefore longer entated. It ends at the base of the lingual flank of stylar cusp B and more obliquely orientated than on M2. The preprotocrista is and does not connect to the premetacrista to form a centrocrista. longer. The metastylar shelf has a weak continuous crest joining The premetacrista ends at the anterior base of the flank of sty- stylar cusp D to the posterolabial corner of the tooth. No poste- lar cusp D. The postmetacrista is the longest crest and runs from rior cingulum is present. the metacone to the posterolabial corner of the tooth. Stylar cusp The morphology of M4 is similar to that of M3 except as fol- A is low, and just anterior to stylar cusp B. The anterior cin- lows (Fig. 9). The anterior cingulum is further developed. The gulum is a continuous shelf from the anterolabial corner of the paracone is lingually level with the metacone of M3. No stylar tooth to the protocone. The protocone is posterolingual to the cusp B is present and no cusp is present on the posterobuccal paracone. The postprotocrista runs posteriorly to the metaconu- edge of the crown. There is no metastylar shelf. The postpro- lar hypocone and curves around the base of the metacone, par- tocrista and postparacrista both terminate at the most posterior allel to the postmetacrista, forming a small posterior cingulum, point on the crown. No metaconular hypocone is present.

Downloaded by [58.165.130.223] at 03:54 06 February 2013 terminating before the posterolabial corner of the tooth. Measurements of the upper dentition of G. amplus are given Morphology of M2 is similar to that of M1 except as follows in Table 3 and those of the lower dentition in Table 4. (Fig. 9). The buccal length and anterior width of the crown are subequal. Stylar cusps B and D are larger and conical in shape. No crest is present between stylar cusps A and B. Stylar cusp A is GALADI ADVERSUS, sp. nov. on the parastylar shelf. The preparacrista is long and joins stylar (Fig. 10) cusp A without connecting to stylar cusp B. The postparacrista and premetacrista terminate at the base of stylar cusp B and sty- Specific Diagnosis—Galadi adversus differs from G. amplus, lar cusp D, respectively. A continuous crest runs from the an- G. grandis,andG. speciosus in the following combination of fea- terior base of stylar cusp D to the posterolabial corner of the tures: infraorbital foramen lies above the posterior root of P3 (as tooth. The protocone and metaconular hypocone lie directly lin- in G. speciosus, above anterior root in G. amplus); postparacrista gual to the paracone and metacone, respectively. The prepro- and premetacrista connect to form a complete centrocrista on tocrista runs anteriorly past the base of the paracone to terminate M3; postparacrista and premetacrista do not connect to form a at the base of anterolabial corner of the tooth, forming a com- complete centrocrista on M1–2; stylar cusp E present on M1–3; plete anterior cingulum. The postprotocristid runs through the stylar cusp B present on M4 (present in G. speciosus but absent metaconular hypocone and bypasses the base of the metacone to in G. amplus); posterior cingulum present on M1 and M2, but terminate on the posterolingual flank of the tooth, forming a pos- not connected to postprotocrista; wider anterior cingulum on M3; terior cingulum. very small metaconular hypocone; and wider metastylar shelf, Morphology of M3 is similar to that of M2 except as follows making the overall upper molar shape more triangular in occlusal (Fig. 9). Stylar cusp A is taller than on M2 but lower than sty- outline. lar cusp B. Stylar cusp B is equal in height to stylar cusp D. Specific Etymology—‘Adversus’, Latin word meaning oppo- Stylar cusp E is present on some specimens. The postprotocrista site, in reference to the centrocrista being complete on M3, and curves and diverges around the metaconular hypocone. The pro- incomplete on M1–2, which is the other way around in Galadi tocone is further lingual than the metaconular hypocone, which is speciosus. TRAVOUILLON ET AL.—FOSSIL BANDICOOTS FROM AUSTRALIA 163

TABLE 3. Measurements (in mm) of the upper dentition of type and referred material of Galadi amplus.

P1 P2 P3 M1 M2 M3 M4 Specimen Riversleigh locality L W L W L W L W L W L W L W QM F23398 Gag Site 3.13 2.99 3.12 3.45 1.91 4.06 QM F23399 Gag Site 2.68 2.13 3.23 2.93.12 3.37 QM F23400 Gag Site 3.47 2.78 3.34 3.48 QM F23401 Gag Site 2.96 2.87 2.94 3.28 1.74 3.99 QM F56069 Gag Site 3.26 3.12 3.04 3.39 QM F56076 Henk’s Hollow Site 3.13.4 QM F56074 Last Minute Site 3.35 2.65 3.07 3.08 1.98 3.89 QMF36308 Wang Site 3.33 3.43 1.87 3.82

Abbreviations: L, anteroposterior length; M,molar;P, premolar; W, lingual-buccal width.

Holotype—QM F16911, maxillary fragment with M1–4 as far as a small metaconular hypocone. A small posterior cin- (Fig. 10). gulum occurs on the posterior flank of the metastylar shelf and Type Locality—Dirk’s Towers Site, Riversleigh WHA, north- runs from level with the metacone to two thirds of the way to the western Queensland, Australia. postmetacrista. Age and Stratigraphy—This site is part of Riversleigh WHA, Morphology of M2 is similar to that of M1 except as follows northwestern Queensland, and contains terrestrial carbonates (Fig. 10). It is shorter wider. Stylar cusps B and D are larger that appear to represent a vadose cave deposits (Arena, 2004). and more conical. Stylar cusp A is on the parastylar shelf and It is part of Faunal Zone B, estimated to be approximately mid- connects to the preparacrista only. A crest runs posterolingually dle Miocene in age (Travouillon et al., 2006). from the tip of stylar cusp B and connects with the postparacrista. The premetacrista connects to a crest at the base of stylar cusp Description D, which continues posteriorly through stylar cusp E and terminates at the posterolabial corner of the tooth. The metaconular Maxilla—The maxilla is poorly preserved. The infraorbital hypocone lies directly lingual to the metacone. foramen is located above the posterior alveolus of P3. The in- Morphology of M3 is similar to that of M2 except as follows fraorbital canal is short and the maxillary foramen is wholly (Fig. 9). Stylar cusp A is taller than on M2, but lower than sty- bounded by the maxilla. A heavily worn fragment of the jugal lar cusps B and D, which are equal in height. No crest runs be- meets the maxilla in a narrow V-shaped maxilla-jugal suture. tween stylar cusp D, stylar cusp E, and the posterolabial corner of Upper Dentition—M1 is longer than wide and its occlusal out- the tooth. The anterior cingulum is further developed. The meta- line is rounded (Fig. 10). Stylar cusp D is the tallest cusp followed conular hypocone is larger. No posterior cingulum is present. by posterolabial corner of the tooth, stylar cusp E, stylar cusp B, The postparacrista connects with the premetacrista to form a metacone, paracone, stylar cusp A, protocone, and metaconular centrocrista. hypocone. From the paracone, the preparacrista runs bucally to- The morphology of M4 is similar to that of M3 except as fol- ward stylar cusp B, then curves and continues to the posterolabial lows (Fig. 9). No metastylar shelf is present. Stylar cusp B is corner of the tooth through the ectoflexus, stylar cusp D, and sty- present but no other cusp or crest is present on the posterobuc- lar cusp E. The postparacrista and premetacrista do not join to cal edge of the crown. No metaconular hypocone is present. The form a centrocrista; instead, the postparacrista ends at the base postprotocrista and postparacrista both terminate at the most of the posterolingual flank of stylar cusp B and the premetacrista posterior point on the crown. ends at the base of the anterolingual flank of stylar cusp D. From Measurements of the upper dentition of G. adversus are given the metacone, the postmetacrista runs to the posterolabial corner in Table 5. of the tooth as the crown’s longest crest. Anterior to stylar cusp Downloaded by [58.165.130.223] at 03:54 06 February 2013 B, the anterolabial corner of the tooth lies below the parastylar RESULTS shelf, with a crest running anteroposteriorly to a small stylar cusp A. The anterior cingulum runs from the anterolabial corner of the In the first analysis of 74 craniodental characters included tooth to the protocone. The postprotocrista runs posteriorly only in our matrix, 71 were parsimony informative. Unconstrained

TABLE 4. Measurements (in mm) of the lower dentition of type and referred material of Galadi amplus.

p1 p2 p3 m1 m2 m3 m4 Specimen Riversleigh locality L W L W L W L AW PW L AW PW L AW PW L AW PW QM F56068 Cleft of Ages Site 3.11 1.58 1.87 3.13 1.79 2.13 3.17 1.99 2.13.45 1.93 1.47 QM F23402 Gag Site 2.48 1.33 2.87 1.84 2.03 3.11 1.96 2.15 QM F23403 Gag Site 2.74 1.49 3.06 1.79 2 3.18 2.05 2.36 3.35 2.04 2.14 QM F23404 Gag Site 2.67 1.11 3.25 2.17 2.26 QM F24226 Gag Site 2.62 1.42.96 2.08 2.16 3.17 1.84 1.57 QM F56075 Gag Site 2.56 1.05 2.81.32.99 1.62 1.86 3.11 1.91.86 QM F56064 Gag Site 3.51 1.98 2.02 3.34 1.86 1.41 QM F56070 Gag Site 2.91.69 2.11 3.31.55 1.28 QM F56072 Henk’s Hollow Site 2.03 0.84 2.75 1.32 2.94 1.77 3.29 1.94 2.43 3.58 2.22.75 3.47 2.21 2.58 3.57 2.04 1.69 QM F56071 Last Minute Site 3.02 1.92 2.33 3.09 2.18 2.26 QM F56073 Last Minute Site 3.36 1.97 2.09 3.47 1.96 1.33

Abbreviations: AW, anterior width; L, anteroposterior length; m,molar;p, premolar; PW, posterior width. 164 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 33, NO. 1, 2013

TABLE 5. Measurements (in mm) of the upper dentition of type and referred material of Galadi adversus.

P1 P2 P3 M1 M2 M3 M4 Specimen Riversleigh locality L W L W L W L W L W L W L W QMF 16911 Dirk’s Towers Site 3.57 2.64 3.53.15 3.65 3.52.08 3.98

Abbreviations: L, anteroposterior length; M,molar;P, premolar; W, lingual-buccal width.

parsimony analysis of the matrix recovered 483 most parsimo- strong support for monophyly of Macrotis species (bootstrap = nious trees (tree length = 315 steps; Fig. 11A). All Galadi and 94%). Yarala taxa are recovered outside the crown group Peramelemor- In the second analysis of 74 craniodental characters included phia, with the relationship of Peroryctes broadbenti unresolved. in our matrix, 72 were parsimony informative. Unconstrained The two Yarala species form a weakly supported clade (bootstrap parsimony analysis of the matrix recovered 15 most parsimo- value of 61%), as does crown group Peramelemorphia (boot- nious trees (tree length = 299 steps; Fig. 11B). All Galadi and strap = 58%). Within the crown group, there is strong support Yarala taxa are recovered outside crown group Peramelemor- for a clade comprising species of Isoodon, Macrotis, Chaeropus, phia and form a sister clade to the crown group. Yarala species and the Pliocene taxa Ischnodon australis (bootstrap = 74%), and form a strong clade (bootstrap = 81%), whereas three of the four Downloaded by [58.165.130.223] at 03:54 06 February 2013

FIGURE 11. Phylogenetic relationship of Galadi amplus, G. grandis, and G. adversus, spp. nov., based on a 74-character craniodental matrix. Fossil and recently extinct taxa are indicated by †. Galadi amplus, G. grandis,andG. adversus, spp. nov., are highlighted in bold. Crown group Peramele- morphia is bracketed. A, Analysis 1 containing all taxa. Strict consensus of 483 most parsimonious trees (tree length = 315; consistency index excluding uninformative characters = 0.3706; retention index = 0.6878) from unconstrained maximum parsimony analysis of the matrix. Numbers above branches represent bootstrap values (100 replicates). Branch lengths are arbitrary. B, Analysis 2 containing all taxa except Galadi adversus, Perameles sobbei, Ischnodon australis, Numbigilga ernielundeliusi,cf. Peroryctes tedfordi,andcf. Peroryctes sp. Strict consensus of 15 most parsimonious trees (tree length = 299; consistency index excluding uninformative characters = 0.3851; retention index = 0.6931) from unconstrained maximum parsimony analysis of the matrix. Numbers above branches represent bootstrap values (100 replicates). Branch lengths are arbitrary. TRAVOUILLON ET AL.—FOSSIL BANDICOOTS FROM AUSTRALIA 165

Galadi taxa (G. adversus was removed in the second analysis) rior tip of nasals = 33.5 mm; rostral width at canine = 13.4 mm), form a weakly supported clade (bootstrap = 57%). Within Gal- suggesting that it would also have fed on relatively larger adi, G. speciosus and G. amplus cluster (bootstrap = 58%). prey. Monophyly of crown group Peramelemorphia including Per- In terms of body mass, Travouillon et al. (2009) calculated the oryctes broadbenti is weakly supported (bootstrap = 58%). estimated body mass of each Galadi species, using the regres- Stronger support (bootstrap = 72%) is found for the crown group sion equations of Myers (2001). Galadi speciosus (formerly ‘Per- excluding Peroryctes broadbenti. Within the crown group, mono- amelemorphia Genus 2 sp. 3’ excluding specimen from Dirk’s phyly of Microperoryctes is recovered (bootstrap = 65%). The Towers Site) ranged between 848 and 987 g (average of 929 g); genus Echymipera is paraphyletic, with strong support (bootstrap G. grandis (formerly ‘Peramelemorphia Genus 2 sp. 2’) ranged = 75%) for a clade that excludes Echymipera rufescens from between 1417 and 1561 g (average of 1488 g); G. amplus (for- other Echymipera taxa and modern Australia genera. A strongly merly ‘Peramelemorphia Genus 2 sp. 1’) ranged between 596 and supported clade containing all taxa from the genera Isoodon, 946 g (average of 763 g); and G. adversus (formerly ‘Peramele- Macrotis, Chaeropus,andPerameles excluding Perameles nasuta morphia Genus 2 sp. 3’ from Dirk’s Towers Site only) was esis recovered. Perameles gunnii and Perameles boungainville form timated at 978 g. Galadi grandis is much larger than the other a clade (bootstrap = 57%). A clade containing Isoodon and three species, but it is found contemporaneously with G. specio- Macrotis is recovered (bootstrap = 69%). There is strong sup- sus in Riversleigh Faunal Zone B sites such as Camel Sputum, port for monophyly of Macrotis (bootstrap = 100%) and Isoodon Mike’s Menagerie, and Wayne’s Wok sites. In modern environ- (bootstrap = 99%). Isoodon obesulus and Isoodon macrourus ments, it is unlikely to find two species from the same genus with- form a well-supported clade (bootstrap = 87%). out having niche partitioning (e.g., size and/or dental morphology of the animal, diet preferences, etc.). Despite having morpholog- DISCUSSION ical similarities, G. grandis and G. speciosus must have occupied different niches based at least on size repartitioning (G. grandis Phylogeny probably took larger prey than G. speciosus) and possibly differ- The phylogenetic relationships proposed here remain unre- ences in diet (G. grandis having more triangular upper molars solved with bootstrap values well under 50% for most clades re- and more complete centrocristae may have been relatively more covered. Removal from the analysis of fossil taxa represented faunivorous than G.speciosus). Galadi speciosus, G. amplus,and only by maxillae or dentaries increased resolution of relation- G. adversus have very similar estimated body masses, but they ships within Peramelemorphia, which is to be expected with no are not contemporaries. G. amplus has a wider body size range characters in common in these taxa; therefore, relationships to than G. speciosus, but this difference may be explained by sexual each other is unresolved, lowering support values (Cobbett et al., dimorphism. Sexual dimorphism is relatively common in bandi- 2007). coots, with extreme dimorphism in Echymipera clara where the Despite this conflict within the crown group Peramelemorphia, male can be more than 5 times larger than the female (Flannery, Galadi species remain outside the crown group in both analyses. 1995). The balance of males versus females in random fossil sam- Although in the first analyses, relationships outside the crown ples may as a result skew the apparent size range for that popu- group were unresolved, the second analysis (after the removal of lation. However, if in this case sampling has fairly sampled the fossil taxa with significant amounts of missing data) shows Gal- population, the size differences could indicate that G. amplus adi species forming a clade sister to the Yarala species (contra was more dimorphic than G. speciosus. It is also possible that Travouillon et al., 2010). This gives sufficient support to conclude G. speciosus, G. amplus,andG. adversus occupied a similar that the taxa described here are in fact from a single genus Gal- niche, and that one species replaced another through time. adi. Cranial and dental features that are shared within the genus Travouillon et al. (2010) noted that fossils of G. speciosus were Galadi are (1) P3 main cusp large and conical, but P3 is narrower relatively uncommon in Riversleigh sites (29 specimens from than M1 (character 5–2); (2) preparacrista connects to the stylar 10 sites) compared with ‘Peramelemorphia Genus 3 sp. 1’ (270 cusp A on both M2 and M3 (character 10–2); and (3) metaconule specimens in 22 sites; see Muirhead, 1994, for species descrip- small but clearly identifiable as a discrete cusp (character 13–1). tion and specimen list; see Archer et al., 2006, for species pres- Our analyses fail to support recent molecular phyloge- ence and reference). G. grandis, G. amplus,andG. adversus are

Downloaded by [58.165.130.223] at 03:54 06 February 2013 nies (Westerman et al., 2001, 2012; Meredith et al., 2008), also quite uncommon (G. grandis—16 specimens from 5 sites; with peroryctines and echymiperines recovered as paraphyletic G. amplus—19 specimens from 5 sites; G. adversus—1 specimen groups, and thylacomyids and chaeropodids nesting within the from 1 site). Carnivores tend to be rarer in the fossil record than peramelids. With increasing fossil taxa and additional mor- herbivores or omnivores (Arita et al., 1990; Carbone et al., 2007), phological characters, this conflict may be resolved. Postcra- which could explain the relative rarity of all species of Galadi. nial elements may be of value in future analyses, although However, the degree of carnivory may differ in the four species. such material is currently known only from the Riversleigh The degree of development of the metaconule, for example, dif- WHA. fers in each of the four species, going from a more triangular tooth (a proxy for relative carnivory) to a more rectangular tooth (a proxy for omnivory). G. adversus has the most triangular up- Ecological Considerations per molars, followed by G. grandis, G. speciosus,andG. amplus, Within the genus Galadi, only G. speciosus and G. amplus are which are increasingly more rectangular. In support of this in- represented by skulls. Travouillon et al. (2010) demonstrated that terpretation, the degree of dental wear in the specimens reflects G. speciosus had a relatively shorter and broader rostrum (skull the same morphological sequence, with G. adversus having the length = 64 mm; rostral length from anterior-most point of or- least dental wear, and G. grandis, G. speciosus,andG. amplus bital rim to anterior tip of nasals = 31.5 mm; rostral width at ca- exhibiting increasingly more wear. High dental wear is usually nine = 12.5 mm) than living bandicoots. Travouillon et al. (2010) correlated with herbivory with plant material being on average concluded that G. speciosus must have been feeding on relatively more abrasive than animal tissue. A recent study (Damuth and larger prey than its contemporary relatives, and therefore was Janis, 2011) demonstrated that increased dental wear in ground- most likely faunivorous. Although the skull of G. amplus is not as based herbivores may also reflect the amount of soil ingested complete that of as G. speciosus, missing the most posterior part while feeding. Most modern bandicoots are omnivorous, feeding of the skull, its rostral dimensions are very similar to G. speciosus on small vertebrates and invertebrates as well as roots and fungi (rostral length from anterior-most point of orbital rim to ante- (Strahan, 1995), but they also have a high rate of dental wear 166 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 33, NO. 1, 2013

FIGURE 12. Diagram showing possible relationships for species in the genus Galadi. A and B, alternative phylogenetic relationships of the four species involved. A, Scenario where Quantum Leap Site (which contains some individuals of G. speciosus) is late Oligocene in age. B,Scenario where Quantum Leap Site is early Miocene in age. C, Patristic hypothesis that G. adversus is an ancestor for G. grandis and G. speciosus and the latter an ancestor for G. amplus. D, Patristic hypothesis that G. adversus was ancestral to G. speciosus and the latter to G. amplus, the three forming an angenetic lineage spanning the early to middle Miocene. Downloaded by [58.165.130.223] at 03:54 06 February 2013 (pers. observ.), which may similarly reflect the amount of soil they Oligocene Quantum Leap Site (Creaser, 1997; Travouillon et al., ingest while consuming ground-based invertebrates and roots. 2006, 2011) would mean it had a longer temporal record than either G. adversus or G. grandis despite being relatively more de- rived (Fig. 12A). However, analyses by Travouillon et al. (2011) Biochronology and Lineage Hypotheses Involving Species suggest that Quantum Leap Site may have too few species (only of Galadi six identified land mammals) to be confidently referred to Faunal As currently known, each of the Galadi species is restricted Zone A. Five other taxa known from this deposit (Namilamadeta temporally. Galadi adversus is only found in Dirk’s Towers Site albivenator, Yalkaparidon coheni, Nambaroo gillespiae, Bulunga- (Faunal Zone B, early Miocene; Travouillon et al., 2006, 2011). maya delicata, and Phalangeridae Genus 2 sp. 1) are present in G. grandis is found in Creaser’s Rampart, Camel Sputum, Mike’s Faunal Zones A and B. Further, Arena (2004) identified Quan- Menagerie, RSO, and Wayne’s Wok sites (Faunal Zone B); tum Leap Site as a cave deposit, as are most Faunal Zone B sites, G. amplus is found in Gag, Henk’s Hollow, Last Minute, Cleft with Faunal Zone A sites tending to be tufa deposits. It is pos- of Ages, and Wang sites (Faunal Zone C, middle Miocene); and sible, therefore, that G. speciosus is restricted to Faunal Zone G. speciosus, with the widest temporal range, is found in Quan- B, and hence G. adversus, which is more plesiomorphic, may tum Leap Site (Faunal Zone A, late Oligocene; but see below), be the first appearance of the genus (Fig. 12B). If this were the as well as Boid, Microsite, Judith’s Horizontalis, Camel Spu- case, Galadi species would be restricted to Faunal Zones B and tum, Mike’s Menagerie, Upper, Neville’s Garden, and Wayne’s C, which both represent rainforest habitats (Travouillon et al., Wok sites (Faunal Zone B). From this, it is clear that three 2009). No Galadi specimens have been found from Encore Site, species of Galadi were contemporaneous in the early Miocene Faunal Zone D (late Miocene), which appears to represent an (see Fig. 12A). However, only G. grandis and G. speciosus have open forest habitat (Travouillon et al., 2009). If Galadi species been found in the same site, and G. adversus is only known are restricted to rainforest habitats, climate change in the middle from Dirk’s Towers Site. Presence of G. speciosus in the late to late Miocene may have led to their local extinction. TRAVOUILLON ET AL.—FOSSIL BANDICOOTS FROM AUSTRALIA 167

As noted above, Galadi adversus is the most plesiomorphic CONCLUSIONS species of Galadi, having a poorly developed metaconule on its The genus Galadi, with four species, is the most diverse fos- upper molars, resulting in more triangular molars, which are sil permelemorphian genus known. Phylogenetically, it is outside more typical of didelphids and dasyurids. Its relationships to all crown group Peramelemorphia and may form a clade with the other species of Galadi remain unclear (see Fig. 11) due to the species of Yarala (Family Yaralidae). Species of Galadi ranged fact that it is only known from one maxilla. Although it appears from an estimated 0.6 to 1.5 kg in size and probably included car- to be the plesiomorphic sister taxon to the other species (Fig. 12A nivorous as well as omnivorous species. As currently understood, and B), there are at least two patristic relationships that are also all species of Galadi were restricted to Riversleigh’s early to mid- possible and not contravened by the phylogenetics, biostratigra- dle Miocene rainforest environments. phy, or prohibitive autapomorphies in potential ancestral taxa: ACKNOWLEDGMENTS 1. G. adversus was an actual ancestor for both G. grandis and G. Support for research at Riversleigh has come from the Aus- speciosus with the latter an ancestor for G. amplus (Fig. 12C). tralian Research Council (grants DP0453262, LP0453664, and 2. G. adversus was an ancestor for G. speciosus and the latter an LP0989969 to M. Archer and S. J. Hand at the University of ancestor for G. amplus (Fig. 12D). New South Wales); XSTRATA Community Partnership Pro- gram (North Queensland); the University of New South Wales; the Queensland National Parks and Wildlife Service; Environ- More fossils specimens of G. adversus will be needed to test these ment Australia; the Queensland Museum; the Australian Mu- hypotheses. seum; the Riversleigh Society Inc.; Outback at Isa; Mount Isa With regard to Megirian et al.’s (2010) interpretation of the City Council; and private supporters including E. Clark, M. Riversleigh biochronology, we did not use it this paper for a num- Beavis, M. Dickson, S. and J. Lavarack, and S. and D. Scott-Orr. ber of reasons. First, Megirian et al. (2010) excluded Riversleigh Assistance in the field has come from many hundreds of volun- from their final seriation analysis because they considered it to teers as well as staff and postgraduate students of the Univer- contain too many anomalous taxa, which they define as species sity of New South Wales (UNSW). We thank the UNSW Palaeo- that span more than one Land Mammal Age (LMA) and were sciences Laboratory for their support. therefore too long-lived. We do not agree with this argument because there are many examples in the literature of long-lived taxa. For example, figure 3 of Prothero (2012) indicates numerous ar- LITERATURE CITED tiodactyls species spanning the Chadronian, Orellan, and Whit- Aplin, K. P., and M. Archer. 1987. Recent advances in marsupial system- neyan NALMAs. atics with a new syncretic classification; pp. xv–lxxii in M. Archer In addition, their scheme is based on the results of a con- (ed.), Possums and Opossums, Studies in Evolution. Surrey Beatty strained seriation analysis. However, Megirian et al. (2010) did and Sons, Sydney, Australia. not give the criterion of the seriation, which would be an indica- Aplin, K. P., K. M. Helgen, and D. Lunde. 2010. 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